Comparison of ecosystem water-use efficiency among Douglas-fir forest, aspen forest and grassland using eddy covariance and carbon isotope techniques

Comparisons were made among Douglas‐fir forest, aspen (broad leaf deciduous) forest and wheatgrass (C₃) grassland for ecosystem‐level water‐use efficiency (WUE). WUE was defined as the ratio of photosynthetic CO₂ assimilation rate and evapotranspiration (ET) rate. The ET data measured by eddy covariance were screened so that they overwhelmingly represented transpiration. The three sites used in this comparison spanned a range of vegetation (plant functional) types and environmental conditions within western Canada. When compared in the relative order Douglas‐fir (located on Vancouver Island, BC), aspen (northern Saskatchewan), grassland (southern Alberta), the sites demonstrated a progressive decline in precipitation and a general increase in maximum air temperature and atmospheric saturation deficit (D_max) during the mid‐summer. The average (±SD) WUE at the grassland site was 2.6±0.7 mmol mol^?1, which was much lower than the average values observed for the two other sites (aspen: 5.4±2.3, Douglas‐fir: 8.1±2.4). The differences in WUE among sites were primarily because of variation in ET. The highest maximum ET rates were approximately 5, 3.2 and 2.7 mm day^?1 for the grassland, aspen and Douglas‐fir sites, respectively. There was a strong negative correlation between WUE and D_max for all sites. We also made seasonal measurements of the carbon isotope ratio of ecosystem respired CO₂ (δ_R) in order to test for the expected correlation between shifts in environmental conditions and changes to the ecosystem‐integrated ratio of leaf intercellular to ambient CO₂ concentration (c_i/c_a). There was a consistent increase in δ_R values in the grassland, aspen forest and Douglas‐fir forest associated with a seasonal reduction in soil moisture. Comparisons were made between WUE measured using eddy covariance with that calculated based on D and δ_R measurements. There was excellent agreement between WUE values calculated using the two techniques. Our δ_R measurements indicated that c_i/c_a values were quite similar among the Douglas‐fir, aspen and grassland sites, despite large variation in environmental conditions among sites. This implied that the shorter‐lived grass species had relatively high c_i/c_a values for the D of their habitat. By contrast, the longer‐lived Douglas‐fir trees were more conservative in water‐use with lower c_i/c_a values relative to their habitat D. This illustrates the interaction between biological and environmental characteristics influencing ecosystem‐level WUE. The strong correlation we observed between the two independent measurements of WUE, indicates that the stable isotope composition of respired CO₂ is a useful ecosystem‐scale tool to help study constraints to photosynthesis and acclimation of ecosystems to environmental stress.     

基于树轮稳定碳同位素的张北杨树防护林退化原因解析

三北防护林是京津冀地区重要的生态屏障,但近十年来张北县4/5的杨树防护林出现退化现象,接近1/3的杨树濒临死亡或枯死,导致杨树防护林生态功能下降.本研究用稳定碳同位素方法追溯研究了退化与未退化杨树年轮中δ¹³C值和内在水分利用效率(WUE_i)的差异,分析导致杨树退化和死亡的原因及其来源.结果表明:相同年龄杨树的直径随着退化程度增加而下降,退化杨树树轮的δ¹³C值变化范围为-25.26‰～-22.97‰,未退化杨树为-26.15‰～-23.50‰,从1997年开始,退化杨树δ¹³C值高于未退化杨树.退化与未退化杨树WUE_i值从1997年开始出现差异但不显著,2002年后其差异达到显著水平,退化与未退化WUE_i的差值ΔWUE_i连续正值可能是退化与未退化杨树产生分化的重要原因之一.退化和未退化杨树WUE_i与降雨量、相对湿度和潜在蒸散量(ET₀)的相关关系均不显著,但与气温和地下水埋深呈极显著的线性关系.1997年极端干旱事件是杨树林退化的起点,随后土地利用方式的改变导致地下水的过度使用,加剧了干旱持续时间和强度,进而加速杨树防护林的退化和死亡.     

树木水分利用效率研究综述

在我国干旱半干旱地区,如何高效地利用极为有限的水资源来合理构建和恢复森林植被,并最大程度地发挥其多种功能,已经成为目前森林生态学研究中的一个热点,而对于树木水分利用效率(WUE)的研究则是其核心和关键内容之一。本文从介绍和剖析树木WUE的概念入手,分别在单叶、个体和群体等三个空间尺度上系统地阐述了WUE概念的内涵及其相应的计算方法,并且对各种方法的优缺点进行了评论。最后,在详细分析国内外树木WUE相关研究进展的基础上,针对我国干旱、半干旱地区水资源短缺和植被恢复与重建之间的矛盾,提出了今后树木WUE的研究发展趋势和应用前景。     

The δ13C of tree rings in full-bark and strip-bark bristlecone pine trees in the White Mountains of California

Dendrochronological work at Sheep Mountain in the White Mountains, CA has demonstrated that bristlecone pine trees in two forms, full-bark and strip-bark, have experienced different cambial growth rates over the past century or longer. The strip-bark trees showed a greater growth increase than the full-bark ones. A calculation of the plant water-use efficiency (W) in response to anthropogenic CO₂ released into the atmosphere shows that W of trees in both forms has increased for the past 200 years. However, there is no significant difference between the two tree forms in the rate of increase in W. This implies at least two possibilities with respect to the CO₂ fertilization effect. First, the biomass in both tree forms might have increased, but carbon distribution among different parts of a tree was different. Second, the biomass may increase without causing any corresponding change in the plant water-use efficiency.     

Modelling environmental controls on ecosystem photosynthesis and the carbon isotope composition of ecosystem-respired CO2 in a coastal Douglas-fir forest

We developed and applied an ecosystem-scale model that calculated leaf CO₂ assimilation, stomatal conductance, chloroplast CO₂ concentration and the carbon isotope composition of carbohydrate formed during photosynthesis separately for sunlit and shaded leaves within multiple canopy layers. The ecosystem photosynthesis model was validated by comparison to leaf-level gas exchange measurements and estimates of ecosystem-scale photosynthesis from eddy covariance measurements made in a coastal Douglas-fir forest on Vancouver Island. A good agreement was also observed between modelled and measured δ ¹³C values of ecosystem-respired CO₂ ( δ _R). The modelled δ _R values showed strong responses to variation in photosynthetic photon flux density (PPFD), air temperature, vapour pressure deficit (VPD) and available soil moisture in a manner consistent with leaf-level studies of photosynthetic ¹³C discrimination. Sensitivity tests were conducted to evaluate the effect of (1) changes in the lag between the time of CO₂ fixation and the conversion of organic matter back to CO₂; (2) shifts in the proportion of autotrophic and heterotrophic respiration; (3) isotope fractionation during respiration; and (4) environmentally induced changes in mesophyll conductance, on modelled δ _R values. Our results indicated that δ _R is a good proxy for canopy-level C _c/ C _a and ¹³C discrimination during photosynthetic gas exchange, and therefore has several applications in ecosystem physiology.     

Carbon isotope discrimination and wood anatomy variations in mixed stands of Quercus robur and Quercus petraea

The two most common oak species in western Europe, Quercus robur and Quercus petraea, display different ecological behaviours, particularly with respect to their responses to drought. The ecophysiological basis of this niche difference is not understood well. Here we test the hypothesis that these two species present distinct water use efficiencies (WUEs), using the carbon isotope discrimination approach. Leaves and 13 dated ring sequences were sampled in 10 pairs of adult trees growing side by side. Carbon isotope composition was measured on cellulose extracts. In addition, relationships between carbon isotope discrimination and wood anatomy were assessed at the tree level. Quercus robur displayed a 1·0‰ larger isotopic discrimination than Q. petraea, and therefore a lower intrinsic WUE (?13%). This interspecific difference of isotopic discrimination was quite stable with time and independent of tree radial growth and climate fluctuations. A strong positive correlation was observed between average tree values of earlywood vessel surface area and ¹³C isotopic discrimination. This correlation was even higher with ¹³C of the 1976 dry year (r = 0·86). These observations led to the hypothesis that hydraulic properties of xylem could exert a constraint on leaf gas exchange, resulting in a larger WUE for individuals with smaller vessel cross‐section area.     

不同生活型绿化植物叶片碳同位素组成的季节特征

通过测定北京地区不同生活型绿化植物叶片的碳同位素组成(δ13C值),从植物种和生活型两个方面研究植物水分利用效率的自然可变性。结果表明:所测定的75种植物(隶属于35科65属)的叶片的δ13C值变幅,春季为-30.7‰--23.4‰,夏季为-31.5‰--25.1‰,秋季为-31.4‰--23.9‰;落叶灌木种间差异不显著(p=0.114),而常绿乔木(p=0.005)、落叶乔木(p0.001)、常绿灌木(p=0.022)、草本植物(p0.001)和藤本植物(p=0.001)的种间差异显著或极显著;同一生活型植物叶片的δ13C季节差异显著,春季叶片的δ13C值显著大于夏秋两季(常绿乔木除外),不同生活型植物叶片的δ13C值在春、夏、秋3个季节差异都达到了极显著水平(春季p=0.001、夏季p0.001、秋季p0.001),且叶片的δ13C值表现出乔木树种灌木树种藤本植物草本植物、常绿植物落叶植物的规律。因此,植物种和生活型均会引起植物叶片δ13C值的变化,但δ13C受生活型变化的影响较大,表明不同生活型植物的水分利用效率具有明显差异。     

长白山阔叶红松林5种主要树种水分利用效率与叶片养分特征

以长白山阔叶红松林中5种优势树种(红松、水曲柳、色木槭、蒙古栎、紫椴)为研究对象,利用稳定碳同位素技术分析了植物的水分利用效率(WUE),测定了叶片碳(C)、氮(N)、磷(P)3种营养元素的含量用以分析植物对养分的利用特征,并综合分析了植物WUE与叶片养分含量之间的关系.结果 表明:冠层不同位置微气象因子的差异导致WU...     

大连市黑松树木水分利用效率的环境响应

以生长于大连城区的黑松为研究对象,建立了1951—2010年间的树木径向生长、树轮稳定碳同位素比率(δ¹³C)和水分利用效率的时间序列,研究了三者的变化特点及其与主要气候因子的关系.结果表明: 1980年以来,黑松树木径向生长有减缓趋势,δ¹³C值降低,但是水分利用效率显著增加(P<0.05).年轮宽度、稳定同位素比率和水分利用效率的变化均受气候因素的影响,并随季节波动:夏季温度与树木径向生长呈负相关,而冬季则呈正相关;6月降水和相对湿度的波动与年轮宽度变化基本呈正相关;3—9月各月温度与δ¹³C和水分利用效率呈弱正相关,其他月份基本呈弱负相关;全年降水和相对湿度分别与δ¹³C和水分利用效率基本呈负相关.快速暖干化的城市气候环境促进了树木水分利用效率的提高.     

Observed and modelled historical trends in the water‐use efficiency of plants and ecosystems

Plant water‐use efficiency (WUE, the carbon gained through photosynthesis per unit of water lost through transpiration) is a tracer of the plant physiological controls on the exchange of water and carbon dioxide between terrestrial ecosystems and the atmosphere. At the leaf level, rising CO₂ concentrations tend to increase carbon uptake (in the absence of other limitations) and to reduce stomatal conductance, both effects leading to an increase in leaf WUE. At the ecosystem level, indirect effects (e.g. increased leaf area index, soil water savings) may amplify or dampen the direct effect of CO₂. Thus, the extent to which changes in leaf WUE translate to changes at the ecosystem scale remains unclear. The differences in the magnitude of increase in leaf versus ecosystem WUE as reported by several studies are much larger than would be expected with current understanding of tree physiology and scaling, indicating unresolved issues. Moreover, current vegetation models produce inconsistent and often unrealistic magnitudes and patterns of variability in leaf and ecosystem WUE, calling for a better assessment of the underlying approaches. Here, we review the causes of variations in observed and modelled historical trends in WUE over the continuum of scales from leaf to ecosystem, including methodological issues, with the aim of elucidating the reasons for discrepancies observed within and across spatial scales. We emphasize that even though physiological responses to changing environmental drivers should be interpreted differently depending on the observational scale, there are large uncertainties in each data set which are often underestimated. Assumptions made by the vegetation models about the main processes influencing WUE strongly impact the modelled historical trends. We provide recommendations for improving long‐term observation‐based estimates of WUE that will better inform the representation of WUE in vegetation models.     

Water use efficiency and carbon isotope composition of plants in a cold desert environment

Summary The effects of the availabilities of water and nitrogen on water use efficiency (WUE) of plants were investigated in a sagebrush steppe. The four species studied wereArtemisia tridentata (shrub),Ceratoides lanata (suffrutescent shrub),Elymus lanceolatus (rhizomatous grass), andElymus elymoides (tussock grass). Water and nitrogen levels were manipulated in a two-by-two factorial design resulting in four treatments: control (no additions), added water, added nitrogen, and added water and nitrogen. One instantaneous and two long-term indicators of WUE were used to testa priori predictions of the ranking of WUE among treatments. The short-term indicator was the instantaneous ratio of assimilation to transpiration (A/E). The long-term measures were 1) the slope of the relationship between conductance to water vapor and maximum assimilation and 2) the carbon isotope composition (¹³C) of plant material. Additional water decreased WUE, whereas additional nitrogen increased WUE. For both A/E and ¹³C, the mean for added nitrogen alone was significantly greater than the mean for added water alone, and means for the control and added water and nitrogen fell in between. This ranking of WUE supported the hypothesis that both water and nitrogen limit plant gas exchange in this semiarid environment. The short- and long-term indicators were in agreement, providing evidence in support of theoretical models concerning the water cost of carbon assimilation.     

Late-stage canopy tree species with extremely low δ13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana

We assessed the daily time‐courses of CO₂ assimilation rate (A), leaf transpiration rate (E), stomatal conductance for water vapour (g_s), leaf water potential ( Ψ _w) and tree transpiration in a wet and a dry season for three late‐stage canopy rainforest tree species in French Guiana differing in leaf carbon isotope composition ( δ ¹³C). The lower sunlit leaf δ ¹³C values found in Virola surinamensis ( ? 29·9‰) and in Diplotropis purpurea ( ? 30·9‰), two light‐demanding species, as compared to Eperua falcata ( ? 28·6‰), a shade‐semi‐tolerant species, were clearly associated with higher maximum g_s values of sunlit leaves in the two former species. These two species were also characterized by a high sensitivity of g_s, sap flow density (Ju) and canopy conductance (g_c) to seasonal soil drought, allowing maintenance of high midday Ψ _w values in the dry season. The data for Diplotropis provided an original picture of increasing midday Ψ _w with increasing soil drought. In Virola, stomata were extremely sensitive to seasonal soil drought, leading to a dramatic decrease in leaf and tree transpiration in the dry season, whereas midday Ψ _w remained close to ? 0·3 MPa. The mechanisms underlying such an extremely high sensitivity of stomata to soil drought remain unknown. In Eperua, g_s of sunlit leaves was non‐responsive to seasonal drought, whereas Ju and g_c were lower in the dry season. This suggests a higher stomatal sensitivity to seasonal drought in shaded leaves than in sunlit ones in this species.     

A retrospective,dual‐isotope approach reveals individual predispositions to winter‐drought induced tree dieback in the southernmost distribution limit of Scots pine

Winter‐drought induced forest diebacks in the low‐latitude margins of species' distribution ranges can provide new insights into the mechanisms (carbon starvation, hydraulic failure) underlying contrasting tree reactions. We analysed a winter‐drought induced dieback at the Scots pine's southern edge through a dual‐isotope approach (Δ¹³C and δ¹⁸O in tree‐ring cellulose). We hypothesized that a differential long‐term performance, mediated by the interaction between CO₂ and climate, determined the fates of individuals during dieback. Declining trees showed a stronger coupling between climate, growth and intrinsic water‐use efficiency (WUEi) than non‐declining individuals that was noticeable for 25 years prior to dieback. The rising stomatal control of water losses with time in declining trees, indicated by negative Δ¹³C‐δ¹⁸O relationships, was likely associated with their native aptitude to grow more and take up more water (suggested by larger tracheid lumen widths) than non‐declining trees and, therefore, to exhibit a greater cavitation risk. Freeze‐thaw episodes occurring in winter 2001 unveiled such physiological differences by triggering dieback in those trees more vulnerable to hydraulic failure. Thus, WUEi tightly modulated growth responses to long‐term warming in declining trees, indicating that co‐occurring individuals were differentially predisposed to winter‐drought mortality. These different performances were unconnected to the depletion of stored carbohydrates.     

The effects of nitrogen stress on the stable carbon isotope composition, productivity and water use efficiency of white spruce (Picea glauca (Moench) Voss) seedlings

A, assimilation rate
a, fractionation against ¹³C for CO₂ diffusion through air
b, net fractionation against ¹³C during CO₂ fixation
C_a, ambient CO₂ concentration
C_c, CO₂ concentration at the chloroplast
C_i, intercellular CO₂ concentration
D, vapour pressure deficit
E_n, needle transpiration rate
E_p, whole plant water use
g_w, leaf internal transfer conductance to CO₂
g_s, stomatal conductance to water vapour
L, projected leaf area
NUE, nitrogen use efficiency
PEP, phosphoenolpyruvate
Rubisco, ribulose-1,5-biphosphate carboxylase
TDR, time domain reflectometry
WUE, water use efficiency
Δ, carbon isotope discrimination
δ¹³C, carbon isotope abundance parameter
δ¹³C_a, carbon isotopic composition of atmospheric CO₂
θ, volumetric soil water content

The effect of nitrogen stress on needle δ¹³C, water-use efficiency (WUE) and biomass production in irrigated and dry land white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings, representing 10 controlled crosses, were planted in the field in individual buried sand-filled cylinders. Two nitrogen treatments were imposed, nitrogen stressed and fertilized. The ranking of δ¹³C of the crosses was maintained across all combinations of water and nitrogen treatments and there was not a significant genetic versus environmental interaction. The positive relationships between needle δ¹³C, WUE and dry matter production demonstrate that it should be possible to use δ¹³C as a surrogate for WUE, and to select for increased WUE without compromising yield, even in nitrogen deficient environments. Nitrogen stressed seedlings had the lowest needle δ¹³C in both irrigated and dry land conditions. There was a positive correlation between needle nitrogen content and δ¹³C that was likely associated with increased photosynthetic capacity. There was some indication that decreased nitrogen supply led to increased stomatal conductance and hence lower WUE. There was a negative correlation between intrinsic water use efficiency and photosynthetic nitrogen use efficiency (NUE). This suggests that white spruce seedlings have the ability to maximize NUE when water becomes limited. There was significant genetic variation in NUE that was maintained across treatments. Our results suggest that in white spruce, there is no detectable effect of anaplerotic carbon fixation and that it is more appropriate to use a value of 29‰ (‘Rubisco only’) for the net discrimination against ¹³C during CO₂ fixation. This leads to excellent correspondence between values of C_i/C_a derived from gas exchange measurements or from δ¹³C.     

Evidence for strong seasonality in the carbon storage and carbon use efficiency of an Amazonian forest

The relative contribution of gross primary production and ecosystem respiration to seasonal changes in the net carbon flux of tropical forests remains poorly quantified by both modelling and field studies. We use data assimilation to combine nine ecological time series from an eastern Amazonian forest, with mass balance constraints from an ecosystem carbon cycle model. The resulting analysis quantifies, with uncertainty estimates, the seasonal changes in the net carbon flux of a tropical rainforest which experiences a pronounced dry season. We show that the carbon accumulation in this forest was four times greater in the dry season than in the wet season and that this was accompanied by a 5% increase in the carbon use efficiency. This seasonal response was caused by a dry season increase in gross primary productivity, in response to radiation and a similar magnitude decrease in heterotrophic respiration, in response to drying soils. The analysis also predicts increased carbon allocation to leaves and wood in the wet season, and greater allocation to fine roots in the dry season. This study demonstrates implementation of seasonal variations in parameters better enables models to simulate observed patterns in data. In particular, we highlight the necessity to simulate the seasonal patterns of heterotrophic respiration to accurately simulate the net carbon flux seasonal tropical forest.     

亚热带地区马尾松树干呼吸碳同位素组成的时间变化特征及其控制因子

作为森林生态系统碳循环的重要组成部分,树干呼吸的时空变异性直接决定着全球碳库对气候变化的响应和反馈。然而,目前关于树干呼吸的时间变化特征,尤其是树干呼吸碳同位素组成(δ¹³C)变化的控制机理还存在很大的不确定性。为探明树干呼吸及其碳同位素的时间变化特征,采用气室法并利用基于光腔衰荡光谱(CRDS)技术组装的Picarro观测系统,于2018年8月—2019年10月对6棵马尾松进行树干呼吸及其同位素的监测,同时测定树干液流速率和树干温度。结果表明:(1)树干呼吸速率及其δ¹³C值均不存在明显的日变化模式,且与树干液流、树干温度相关性不显著;(2)树干呼吸速率季节变化趋势明显,变化范围为0.08～1.61μmol·m^-2·s^-1,且树干温度解释了树干呼吸速率季节变化的82%;(3)树干呼吸δ¹³C呈现先升高后下降的季节变化模式,非生长季的平均δ¹³C为-23.25‰±0.91‰,显著高于生长季的-27.93‰±0.25‰。研究表明,树干呼吸及其碳同位素组成在不...